garciaberthou_etal_gherardi_ed_book.doc
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Non-indigenous animal species naturalized in Iberian inland waters Emili Garcı́a-Berthou, Dani Boix, and Miguel Clavero INTRODUCTION Invasions by human-introduced non-indigenous species (NIS) are one of the main threats to biodiversity and a driving force of global change (Vitousek et al. 1997, Mack et al. 2000, Clavero and Garcı́a-Berthou 2005). The Iberian Peninsula (IP) is a hotspot of biodiversity (Médail and Quézel 1999) and a knowledge of the invasive species inhabiting it is essential for conservation and environmental management. Naturalized vertebrates and plants in the IP have received considerable attention (see e.g. Vilà et al. 2001, Pleguezuelos 2002, Sobrino et al. 2002, Lloret et al. 2004, Alcaraz et al. 2005), but its invasive invertebrates are very poorly known. Although there are many records of some invertebrate invasive species, particularly crustaceans, there are very few available reviews of selected taxa of invertebrate invaders in the IP (e.g. Espadaler and Collingwood 2001). The aim of this chapter is to review the animal species naturalized in Iberian inland waters, including vertebrates and free-living and parasitic invertebrates. As usual, the taxonomy and biogeography of vertebrate species are much better known than for invertebrates, so our data for invertebrates should be regarded as a preliminary check-list. Similarly, the parasites of non-commercial aquatic species are poorly studied and the data in the IP mostly come from studies of the eel, Anguilla anguilla (Linnaeus), thus certainly underestimating the range of introduced parasites (Blanc 1997, 2001). We feel, however, that it is important to provide such a . first check-list because many of the invertebrates involved are nowadays common in the IP and for many of them it is largely unknown even by biologists that they are not indigenous to the IP. Increasing the awareness on the introduced status and current distribution of these species is essential to reduce their spread and impact. We compiled animal species cited (by March 2006) as currently naturalized in Iberian inland waters from the scientific literature and unpublished Spanish Ph.D. theses (http://teseo.mec.es/teseo/). We included species from estuaries and saline coastal lagoons but excluded purely marine taxa and terrestrial animal species not strictly linked to aquatic ecosystems. We list invertebrate and vertebrate species introduced by humans and currently naturalized, i.e. species that reproduce and sustain populations in the wild without human intervention (see e.g. Richardson et al. 2000, Pyšek et al. 2004). A few uncertain cases are listed in a separate table. The introduced origin of parasite invertebrates is particularly uncertain but we followed Blanc (1997, 2001), who has recently provided a comprehensive list of aquatic parasites introduced to Europe, together with their native distribution. NATURALIZED ANIMALS IN IBERIAN INLAND WATERS The invertebrate and vertebrate species naturalized in Iberian inland waters are listed in Tables 1 and 2, respectively. A few cases, for which it is not clear whether the species is indigenous to the IP or whether they have established, are listed in Table 3. We found 45 invertebrate and 28 vertebrate species certainly naturalized at present in Iberian inland waters. Among the 45 invertebrates, 12 were parasites (mostly Platyhelminthes flatworms), mainly of freshwater fish and introduced to Europe from Asia with common carp (Cyprinus carpio), goldfish (Carassius auratus), or Japanese eel (Anguilla japonica Temminck and Schlegel) (see Blanc 1997, 2001); several of the parasites have now been recorded on fish species indigenous or endemic to the IP (see references in Table 1). The remaining 33 invertebrates were free-living species, mostly crustaceans (18 species) or molluscs (6 species). Most of the 28 vertebrates were fish (23 species), and there was no aquatic bird naturalized and only one amphibian and one reptilian species. The continent of origin was significantly different between vertebrates and invertebrates (independence test; x 2 ¼ 37:1, df ¼ 7, P < 0.0005) because most naturalized vertebrates were native to the rest of Europe (43% of the 28 species) or North America (29%), origins that in turn were rare among invertebrates (0 and 12%, respectively), which predominantly came from Asia (38%). There was no significant variation in origin between free-living and parasitic invertebrates (x 2 ¼ 6:5, df ¼ 5, P ¼ 0.26) or between crustaceans and molluscs (x 2 ¼ 3:8, df ¼ 5, P ¼ 0.59). Table 1 Non-indigenous invertebrate species naturalized in inland waters of the lberian Peninsula (IP). When a species is probably an old introduction, but the introduction date is largely unknown, a question mark is given as the first record date. The habitats for nonparasite species are coded as: 1, streams and rivers (excluding estuaries); 2, lakes and reservoirs; 3, ponds and pools; 4, rice fields; and 5, estuarine or saline waters. Indigenous distribution CNIDARIA Cordylophora caspia (Pallas) Craspedacusta sowerbyi Lankester Haliplanella lineata (Verrill) Ponto–Caspian Asia Pacific coast First record Habitat/ in the IP Parasite References for the IP 2001 1968 1996 1,2,5 1,2 5 PLATYHELMINTHES TURBELLARIA Dugesia tigrina (Girard) North America ? 1,2 PLATYHELMINTHES MONOGENEA Dactylogyrus anchoratus (Dujardin) Asia Gyrodactylus cyprini Diarova Asia Gyrodactylus katharineri Malmberg Asia ? ? ? Gyrodactylus salaris Malmberg Pseudodactylogyrus anguillae (Yin and Sproston) Baltic Asia and Australia ? ? Pseudodactylogyrus bini (Kikuchi) Asia and Australia ? Other references Schuchert 2004 Escot et al. 2003, Solà 2004 Margalef 1974, Ferreira 1985 Gollasch and RiemannCuesta et al. 1996 Zürneck 1996 Baguñ à et al. 1980 parasite Sánchez Suá rez 2002 parasite Lacasa Millán 1992 parasite Gutiérrez Galindo and Lacasa Millán 1999 parasite Bakke et al. 2002 parasite San Martı́n Outeiral 2004, Gómez-Juaristi and Salvador 2006 parasite Orts Muñ oz 1993, GómezJuaristi and Salvador 2006 Young and Reynoldson 1999 Blanc 1997, 2001 Blanc 1997, 2001 Blanc 1997, 2001 Animal NIS in Iberian inland waters GROUP Species Blanc 1997, 2001 Blanc 1997, 2001 Blanc 1997, 2001 125 126 Table 1 Continued. GROUP Species Ponto–Caspian? First record Habitat/ in the IP Parasite References for the IP Other references 2004 parasite Peribá ñ ez et al. 2006 Asia ? parasite Lacasa Millán 1992 Blanc 1997, 2001 Asia ? parasite Gallastegi et al. 2002, Maillo et al. 2005 Blanc 1997, 2001 Asia North America New Zealand 1979 1845 1951 2,4 1,2,4 1,5 Brown et al. 1998 Anderson 2005 Vidal-Abarca and Suá rez 1985 Anderson 2003 Vidal-Abarca and Suá rez 1985 Paavola et al. 2005 Asia, Africa, and Australia Ponto–Caspian 1981 1,3,5 1880 1,2 Mytilopsis leucophaeta (Conrad) ANNELIDA Branchiura sowerbyi Beddard Gulf of Mexico 1993 1,2,5 Vidal-Abarca and Suá rez 1985, McMahon 2000 Escot et al. 2003 Vidal-Abarca and Suá rez 1985, Altaba et al. 2001 Escot et al. 2003 Laine et al. 2006 Asia 1970s 1,2,5 Prat 1980 Xironogiton victoriensis Gelder and Hall North America 1996 Dreissena polymorpha (Pallas) parasite Gelder 1999 Brinkhurst and Jamieson 1971 Ohtaka et al. 2005 Emili Garcı́a-Berthou et al. PLATYHELMINTHES TREMATODA Phyllodistomum folium (Olfers, 1816) PLATYHELMINTHES CESTODA Bothriocephalus acheilognathi Yamaguti [¼ Bothriocephalus opsarichthydis (Yeh)] NEMATODA Anguillicola crassus Kuwahara, Niimi and Itagaki MOLLUSCA GASTROPODA Gyraulus chinensis (Dunker) Physella acuta (Draparnaud) Potamopyrgus antipodarum (Gray) [¼ P. jenkinsi (Smith)] MOLLUSCA BIVALVIA Corbicula fluminea (Müller) Indigenous distribution Ficopomatus enigmaticus (Fauvel) (¼ Mercierella enigmatica Fauvel) CRUSTACEA BRANCHIOPODA Artemia franciscana (Kellog) 1924 5 Rioja 1924, Fischer-Piette 1951 North, Central, and South America Eurasia 1980s 5 Amat et al. 2005 1990s 4 Martinoy et al. 2006 Leoni et al. 1999 Asia 1986 3,4 Rossi et al. 2003 1996 4 Cypris sp. (¼ Cypris subglobosa Sowerby) Isocypris beauchampi (Paris) Australia and New Zealand America, Africa, and Asia? Africa Forés et al. 1986, Baltaná s et al. 1996 Baltaná s et al. 1996 1986 2,4 Whatley et al. 2003 1976 2,4 Stenocypris major (Baird) Asia 1986 3,4 Strandesia vavrai (Müller) Africa 1983 4 Strandesia vinciguerrae (Masi) Africa 1986 4 Tanycypris sp. Asia 1988 3,4 Forés et al. 1986, Baltaná s et al. 1996 Armengol 1976, Baltaná s et al. 1996 Forés et al. 1986, Baltaná s et al. 1996 Paulo and Moutinho 1983, Baltaná s et al. 1996 Forés et al. 1986, Baltaná s et al. 1996 Forés 1988, Baltaná s et al. 1996 North and South America Asia 1990s 5 Wlassicsia pannonica Daday CRUSTACEA OSTRACODA Dolerocypris sinensis Sars Ilyodromus viridulus (Brady) CRUSTACEA COPEPODA Acartia tonsa Dana Lernaea cyprinacea Linnaeus Rossi et al. 2003 Rossi et al. 2003 Rossi et al. 2003 Rossi et al. 2003 Animal NIS in Iberian inland waters Indian Ocean Sobral 1985, Frisch et al. 2005 Blanc 1997, 2001 127 parasite Moreno et al. 1986, GutiérrezGalindo and Lacasa-Millán 2005 128 Table 1 Continued. Indigenous distribution CRUSTACEA BRANCHIURA Argulus japonicus Thiele CRUSTACEA MYSIDACEA Synidotea laticauda Benedict CRUSTACEA DECAPODA Cherax destructor Clark Eriocheir sinensis (Milne Edwards) Pacifastacus leniusculus (Dana) Palaemon macrodactylus Rathbun Procambarus clarkii (Girard) Rhithropanopeus harrisii (Gould) INSECTA Aedes albopictus (Skuse) Stenopelmus rufinasus Gyllenhal Trichocorixa verticalis (Fieber) First record in the IP Habitat/ Parasite Asia 1921 parasite North Pacific and South Atlantic coasts 1996 5 Australia Asia North America Asia North America North-west Atlantic 1983 ? 1974 1999 1974 1991 5 5 1 5 1,2,4,5 5 Gutiérrez-Yurrita et al. 1999 Cuesta et al. 2006 Habsburgo-Lorena 1978 Cuesta et al. 2004 Habsburgo-Lorena 1978 Cuesta et al. 1991 Asia Asia Atlantic coast of America 2004 2003 1997 3 1,2,4,5 3,5 Aranda et al. 2006 Dana and Viva 2006 Günther 2004, Sala and Boix 2005 References for the IP Other references http://www.fauna-iberica. mncn.csic.es/ Blanc 1997, 2001 Cuesta et al. 1996 Eritja et al. 2005 Hutchinson 1931 Emili Garcı́a-Berthou et al. GROUP Species Animal NIS in Iberian inland waters 129 Table 2 Non-indigenous vertebrate species naturalized in inland waters of the Iberian Peninsula (IP). The habitats are coded as: 1, rivers (excluding estuaries); 2, lakes and reservoirs; 3, rice fields; and 4, estuarine or saline waters. GROUP Species PISCES Abramis bjoerkna (Linnaeus) Abramis brama (Linnaeus) Alburnus alburnus (Linnaeus) Ameiurus melas (Rafinesque) Carassius auratus Linnaeus Cobitis bilineata Canestrini Cyprinus carpio Linnaeus Esox lucius Linnaeus Fundulus heteroclitus (Linnaeus) Gambusia holbrooki (Girard) Herichthys facetum (Jenyns) Hucho hucho (Linnaeus) Lepomis gibbosus (Linnaeus) Micropterus salmoides (Lacepède) Oncorhynchus mykiss (Walbaum) Perca fluviatilis Linnaeus Poecilia reticulata Peters Pseudorasbora parva (Temminck and Schlegel) Rutilus rutilus (Linnaeus) Salvelinus fontinalis (Mitchill) Sander lucioperca (Linnaeus) Scardinius erythrophthalmus (Linnaeus) Silurus glanis L. AMPHIBIA ANURA Discoglossus pictus Otth REPTILIA CHELONIA Trachemys scripta (Schoepf ) MAMMALIA Mustela vison Schreber Myocastor coypus Molina Ondatra zibethicus (Linnaeus) Indigenous distribution First record in the IP Habitat References for the IP Europe 1995 Europe 2004 Europe 1992 North America 1910 Asia 17th century Europe 2002 Eurasia 17th century Europe 1949 North America 1970 North America South America Europe North America North America 1920 1985 1970 1910 1955 1,2 2 1,2 1,2 1,2,3 1 1,2 1,2 4 1,2,3,4 1,2 1 1,2 1,2 Doadrio 2002 Benejam et al. 2005 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio Doadrio Doadrio Doadrio Doadrio 2002 2002 2002 2002 2002 North America 19th century Europe 1975 South America 2000 Asia 2001 1,2 1,2 1,4 1 Europe 1910 North America 19th century Europe 1975 Europe 1910 1,2 1,2 1,2 1,2 Doadrio 2002 Doadrio 2002 Doadrio 2002 Caiola and Sostoa 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Doadrio 2002 Europe 1974 1,2 Doadrio 2002 Africa 1900 1 America 1985 1,2,3,4 Pleguezuelos 2002 North America 1978 South America North America 1970 2002 1,2,3,4 Ruiz-Olmo et al. 1997, Palomo and Gisbert 2002 1 Palomo and Gisbert 2002 1 Elosegi 2004 Pleguezuelos 2002 130 Emili Garcı́a-Berthou et al. Table 3 Animal species possibly introduced to inland waters of the Iberian Peninsula (IP), but with uncertain status. Some species are cryptogenic (Carlton 1996), i.e. it is very difficult to know whether they are indigenous or introduced; the other species have been reported in the wild but it is uncertain whether they have established permanent populations (naturalized). GROUP Species MOLLUSCA GASTROPODA Ferrissia wautieri (Mirolli) [¼ F. clessiniana (Jickeli)] CRUSTACEA DECAPODA Austropotamobius italicus (Faxon) / Austropotamobius pallipes (Lereboullet) PISCES Acipenser baeri Brandt Aphanius fasciatus (Valenciennes) Ctenopharyngodon idella (Valenciennes) Ictalurus punctatus (Rafinesque) Oncorhynchus kisutch (Walbaum) Tinca tinca (Linnaeus) AMPHIBIA Bufo mauritanicus Schlegel Rana catesbeiana Shaw Rana ridibunda, Rana kl. esculenta, Rana lessonae REPTILIA CHELONIA Pelodiscus sinensis (Wiegmann) AVES Aix galericulata (Linnaeus) Anser erythropus (Linnaeus) Branta canadensis (Linnaeus) Oxyura jamaicensis (Gmelin) MAMMALIA Castor fiber Linnaeus Possibly indigenous to the IP Uncertain establishment yes no yes no Grandjean et al. 2001 no no no yes yes yes no no yes yes yes no Elvira and Almodóvar 2001 Doadrio 2002 J. M. Queral 2005, personal communication Doadrio 2002 Doadrio 2002 Doadrio 2002 no no no yes yes yes Pleguezuelos 2002 Pleguezuelos 2002 Arano et al. 1995, Garcı́a-Parı́s et al. 2004 no yes Pleguezuelos 2002 no no no no yes yes yes yes GAE GAE GAE GAE yes yes Ceña et al. 2004 References Anderson 2005 2006 2006 2006 2006 The main habitat also differed between vertebrates and invertebrates (independence test; x 2 ¼ 22:8, df ¼ 4, P < 0.0005), because the former were mostly present in streams and rivers (26 of the 28 species were present in streams and rivers) or lakes and reservoirs, whereas several invertebrates were only present in estuaries/saline waters (e.g. several decapod crustaceans introduced into the Guadalquivir River through ballast water) or in rice fields (namely ostracods). Animal NIS in Iberian inland waters 131 The mechanism of introduction is obviously also different for invertebrates and vertebrates, because most of the former are accidental introductions (e.g. Asian ostracods in rice fields, ballast water, etc.), whereas most fish species have been introduced intentionally (nowadays illegally). Therefore, naturalized vertebrates and invertebrates showed opposite patterns, with the former (mostly fish) intentionally introduced from the rest of Europe or North America to Iberian streams and reservoirs and most invertebrates originating from Asia and accidentally introduced to estuaries or rice fields. UNCERTAIN CASES We found four species for which it is uncertain whether the species is indigenous to the IP and 13 species that they may not have established (Table 3). An interesting case illustrating both the lack of knowledge on invasive species and the power of modern genetic techniques is the crayfish of the Austropotamobius pallipes species complex. Until the 1980s the populations in the IP were generally regarded as an endemic species or subspecies in strong decline due to the introduction of the oomycete Aphanomyces astaci Schikora with North American crayfish (Martı́nez et al. 2003). Grandjean et al. (2000) showed that two species (A. pallipes and Austropotamobius italicus) could be distinguished within the species complex and that Spanish populations were very close to some Italian populations, so they might be of anthropogenic origin, as already proposed by Albrecht (1983), and should be regarded as A. italicus. Grandjean et al. (2001) demonstrated a drastic bottleneck in Spanish populations but discussed several potential mechanisms alternative to the hypothesis of introduction by humans. With further genetic analyses, Trontelj et al. (2005) supported the anthropogenic origin for the Spanish populations but did not find unequivocal separation between A. pallipes and A. italicus (but see also Schulz and Grandjean 2005). These genetic techniques might also prove useful for tench [Tinca tinca (Linnaeus)] in the IP. Tench is indigenous to many parts of Europe but considered introduced into Italy (Bianco 1998) and Portugal (Almaça 1995). This latter country shares its largest river basins (Duero, Tajo, and Guadiana rivers) with Spain. There are doubts about its indigenous status in Spain (Doadrio 2002). In fact, Gó mez Caruana and Dı́az Luna (1991) considered it introduced into the IP around the 17th century. There are records of tench stocking by monks in Spanish and Portuguese ponds several centuries ago (Almaça 1995, Garcı́a-Berthou and Moreno-Amich 2000). As far as we know, no phylogeographic study on tench has been performed, in contrast to many other European cyprinids, although they could be most helpful in clarifying its native distribution. A similar, more solved example of ‘‘cryptogenic’’ species (see Carlton 1996) is the case of the freshwater snail Physella acuta (Draparnaud). This species was first described from Europe (Drapanaud, 1805), namely from the River 132 Emili Garcı́a-Berthou et al. Garonne, near Bordeaux (France). This species is widely distributed in the IP and the rest of western Europe (Vidal-Abarca and Suá rez 1985) and inhabits all types of fresh waters. It has been generally regarded as indigenous to continental Europe (Haas 1929, Germain 1930, Macan and Cooper 1977, Girod et al. 1980, Vidal-Abarca and Suá rez 1985) and its presence in North America was not reported until the 1990s (Wu et al. 1997). Nowadays, three types of evidence indicate, however, that P. acuta is indigenous to North America and not to Europe: (i) the lack of records of Physella shells from European sediments older than the 18th century (Lozek 1964); (ii) recent studies using internal morphology comparisons (Anderson 2003) and reproductive isolation experiments (Dillon et al. 2002) showing that at least one Physella species from North America [Physella heterostropha (Say)] is actually P. acuta; and (iii) some historical data of the cotton trade between France and the United States in the 18th century that could explain the arrival of this species to the River Garonne, where it was first observed (Anderson 2003). The case of P. acuta illustrates the importance of historical data and the fossil record as tools for the identification of old introductions by man. Fossil records have been very helpful to establish the introduced nature of ostracods and suggest that dispersal by man of many other invertebrates is very old and has been generally neglected (McKenzie and Moroni 1986, Rossi et al. 2003). The other group of species in Table 3 are species that have been reported in the wild but it is uncertain whether they have established. There are several other NIS that have been recorded in the wild (see e.g. Elvira and Almodó var 2001, Pleguezuelos 2002) but have certainly not established permanent populations. ECOLOGICAL IMPACT The ecological impact of most of these NIS is largely unknown with a few exceptions. The red swamp crayfish, Procambarus clarkii, has altered the functioning and structure of many aquatic ecosystems in the IP reducing macrophytes and associated species, among other impacts (Geiger et al. 2005, Rodrı́guez et al. 2005, Chapter 28). The eastern mosquitofish (Gambusia holbrooki) has been experimentally demonstrated to affect endemic cyprinodontiform fishes [Aphanius iberus (Valenciennes) and Valencia hispanica (Valenciennes)] by resource and interference competition (Rincón et al. 2002). The zebra mussel (Dreissena polymorpha) is one of the best known invasive species and, although it is a very old introduction into Portugal, only recently has it been introduced to Spain through the Ebro River, where it is widespread nowadays and might affect the endangered giant pearl mussel, Margaritifera auricularia Spengler (Altaba et al. 2001). The zebra mussel is still not widespread in the IP, but it will probably be fostered by the illegal, poorly controlled introduction and translocations of fish that are still very frequent. Animal NIS in Iberian inland waters 133 The polychaete Ficopomatus enigmaticus (Fauvel) is very abundant in some Spanish coastal lagoons and probably profoundly affects its ecosystem functioning because it builds large reef-like aggregates (Schwindt and Iribarne 1998). Many piscivorous fish have been introduced into the IP and some unique ecosystems such as Lake Banyoles have been profoundly altered and are nowadays completely dominated by NIS (Garcı́a-Berthou and Moreno-Amich 2000). The distribution, abundance, and impact of introduced parasites in the IP is largely unknown but some species such as Lernaea cyprinacea are widespread (Moreno et al. 1986, Gutiérrez-Galindo and Lacasa-Millá n 2005) and several of them have now been recorded on endemic fish species (see references in Table 1). The swimbladder nematode Anguillicola crassus, which was transferred from its indigenous host (the Japanese eel, A. japonica) to the European eel (A. anguilla), can severely impair swimbladder function (and thus possibly spawning migration) and has caused mortalities in both farmed and wild populations in the presence of other stressors (Kirk 2003). Similarly to the case of crayfish plague, Gozlan et al. (2005) have recently shown that the topmouth gudgeon, Pseudorasbora parva, an Asiatic cyprinid highly invasive in Europe and recently introduced to the IP, carries a pathogen that strongly affects indigenous cyprinids. Given the enormous impact of the few well-investigated invasive species, the considerable number of introduced species, and the presence in the IP of many endemic species of plants (Médail and Quézel 1999), freshwater fish (Doadrio 2002), and amphibians (Pleguezuelos et al. 2002), the overall potential impact of these naturalized species is enormous and should be urgently investigated. The room for management and educational improvement by public administrations to prevent further introductions and translocations and to reduce the spread of invasive species is even larger. We hope this paper will contribute to the improved understanding and control of invasive species in European waters. ACKNOWLEDGEMENTS We thank J.F. Gutiérrez-Galindo, F. Padró s, and anonymous reviewers for helpful comments. 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